Biotic interactions affect the elevational ranges of high‐latitude plant species

Ecological theory suggests that positive plant–plant interactions can extend species distributions into areas that would otherwise be unfavourable. However, few studies have tested this hypothesis, and none have explicitly examined the associated prediction that inter‐specific interactions between plants may broaden species altitudinal distributions. Here we test this prediction, using fine‐scale species distribution data for 156 bryophytes, lichens and vascular plants spanning a 900 m elevational gradient in north‐western Finland and Norway, analysed with a niche modelling approach. Species altitudinal ranges of all three groups of plants were more accurately predicted when including the cover of any of the 24 most wide‐spread and abundant species (‘dominants’) than when using abiotic variables alone, emphasizing the importance of including relevant biotic predictors in species distribution models. Half of the models showed that species had very low probabilities of occurrence under high cover of dominants, suggesting a strong negative impact of dominant species. Similarly, for species that are predicted to occur irrespective of dominant species cover, 62% of models showed narrower species altitudinal distributions when occurring under high dominant cover, with contractions of species’ lower and upper elevational limits being common. Nonetheless, high cover of dominant species was associated with upslope range extension in 43 species, and a net range expansion in nearly 10% of all models. Species distributional responses to dominants were only weakly related to species traits, with larger range contractions associated with arctic‐alpine dominants. Therefore, dominant species appear to exert a strong influence on the elevational distribution of other species in high latitude environments.     

Influences of interpolation of species ranges on elevational species richness gradients

Interpolation of species ranges has been a common approach to compensate for the unevenness or incompleteness in sampling effort in studies of geographic species richness gradients. However, potential biases introduced by this estimation method remain unclear. Here, we presented an explicit examination of the influences of one‐dimensional interpolation on elevational species richness gradients, and discussed potential causes and processes of these influences. We conducted intensive surveys of birds along the elevational gradients of the Ailao Mountains, southwestern China, and compared richness patterns based on interpolation with raw data as well as estimated data from rarefaction and Chao1 non‐parametric estimator; we also compared results of multiple linear regressions and hierarchical partitioning analyses explaining these four measures of richness. Actual evapotranspiration (AET) and the mid‐domain effect (MDE) were highly correlated and separately provided a good potential explanation for the unimodal richness pattern in the Ailao Mountains, with modifying and suppressive effects of other variables such as area. Interpolation consistently and significantly increased the effects of AET/MDE, while it reduced contributions of area and human disturbance. Our results demonstrated that while compensating for biases in sampling effort, interpolation may also spuriously fill genuine distribution gaps, and tend to underestimate the effects of the non‐monotonic or discontinuous influencing factors that are responsible for these gaps, and overestimate the effects of other factors actually suppressed by these factors. These influences were most strong for species with relatively medium elevational ranges. We conclude that at the regional scale, interpolation method is a potential source of bias in identifying and explaining species richness gradients and should be used with careful consideration. It may be advantageous to adopt other robust estimation methods besides interpolation to gain a more accurate assessment of species richness and a more objective understanding of their underlying mechanisms.     

Fern species richness along a central Himalayan elevational gradient, Nepal

Aim The study explores fern species richness patterns along a central Himalayan elevational gradient (100–4800 m a.s.l.) and evaluates factors influencing the spatial increase and decrease of fern richness. Location The Himalayas stretch from west to east by 20°, i.e. 75–95° east, and Nepal is located from 80 to 88° east in this range. Methods We used published data of the distribution of ferns and fern allies to interpolate species elevational ranges. Defining species presence between upper and lower elevation limit is the basis for richness estimates. The richness pattern was regressed against the total number of rainy days, and gradients that are linearly related to elevation, such as length of the growing season, potential evapotranspiration (PET, energy), and a moisture index (MI = PET/mean annual rainfall). The regressions were performed by generalized linear models. Results A unimodal relationship between species richness and elevation was observed, with maximum species richness at 2000 m. Fern richness has a unimodal response along the energy gradients, and a linear response with moisture gradients. Main conclusions The study confirms the importance of moisture on fern distributions as the peak coincides spatially with climatic factors that enhance moisture levels; the maximum number of rainy days and the cloud zone. Energy‐related variables probably control species richness directly at higher elevations but at the lower end the effect is more probably related to moisture.     

An elevational shift in butterfly species richness and composition accompanying recent climate change

The geographic ranges of many species have shifted polewards and uphill in elevation associated with climate warming, leading to increases in species richness at high latitudes and elevations. However, few studies have addressed community‐level responses to climate change across the entire elevational gradients of mountain ranges, or at warm lower latitudes where ecological diversity is expected to decline. Here, we show uphill shifts in butterfly species richness and composition in the Sierra de Guadarrama (central Spain) between 1967–1973 and 2004–2005. Butterfly communities with comparable species compositions shifted uphill by 293 m (± SE 26), consistent with an upward shift of approximately 225 m in mean annual isotherms. Species richness had a humped relationship with elevation, but declined between surveys, particularly at low elevations. Changes to species richness and composition primarily reflect the loss from lower elevations of species whose regional distributions are restricted to the mountains. The few colonizations by specialist low‐elevation species failed to compensate for the loss of high‐elevation species, because there are few low‐elevation species in the region and the habitat requirements of some of these prevent them from colonizing the mountain range. As a result, we estimated a net decline in species richness in approximately 90% of the region, and increasing community domination by widespread species. The results suggest that climate warming, combined with habitat loss and other drivers of biological change, could lead to significant losses in ecological diversity in mountains and other regions where species encounter their lower latitudinal‐range margins.     

Changes to the elevational limits and extent of species ranges associated with climate change

The first expected symptoms of a climate change‐generated biodiversity crisis are range contractions and extinctions at lower elevational and latitudinal limits to species distributions. However, whilst range expansions at high elevations and latitudes have been widely documented, there has been surprisingly little evidence for contractions at warm margins. We show that lower elevational limits for 16 butterfly species in central Spain have risen on average by 212 m (± SE 60) in 30 years, accompanying a 1.3 °C rise (equivalent to c. 225 m) in mean annual temperature. These elevational shifts signify an average reduction in habitable area by one‐third, with losses of 50–80% projected for the coming century, given maintenance of the species thermal associations. The results suggest that many species have already suffered climate‐mediated habitat losses that may threaten their long‐term chances of survival.     

植物物种多样性的垂直分布格局

生物多样性沿环境梯度的变化趋势是生物多样性研究的一个重要议题,而海拔梯度包含了多种环境因子的梯度效应,因此研究生物多样性的海拔梯度格局对于揭示生物多样性的环境梯度变化规律具有重要意义。在不同的研究尺度,植物多样性沿海拔梯度具有不同的分布格局,而形成这种格局的因素有很大差异。本文从α多样性,β多样性和γ多样性三个尺度总结了植物物种多样性沿海拔梯度分布格局及其环境解释。α多样性沿海拔梯度的分布格局在不同生活型的物种之间差异很大,但对于木本植物来说,虽然也存在其他格局,但α多样性随海拔升高而降低是被广泛接受的一种格局。在一般情况下,β多样性随着海拔的升高而降低,并且对于不同生活型的物种,β多样性沿海拔梯度具有相似的分布格局。γ多样性沿海拔梯度具有两种分布格局：偏峰分布格局和显著的负相关格局;特有物种数往往随着海拔的升高而减少,而特有度则随着海拔的升高而增加。     

Mountain roads shift native and non‐native plant species' ranges

Roads are known to act as corridors for dispersal of plant species. With their variable microclimate, role as corridors for species movement and reoccurring disturbance events, they show several characteristics that might influence range dynamics of both native and non‐native species. Previous research on plant species ranges in mountains however seldom included the effects of roads. To study how ranges of native and non‐native species differ between roads and adjacent vegetation, we used a global dataset of plant species composition along mountain roads. We compared average elevation and range width of species, and used generalized linear mixed models (GLMMs) to compile their range optimum and amplitude. We then explored differences between roadside and adjacent plots based on a species’ origin (native vs non‐native) and nitrogen and temperature affinity. Most non‐native species had on average higher elevational ranges and broader amplitudes in roadsides. Higher optima for non‐native species were associated with high nitrogen and temperature affinity. While lowland native species showed patterns comparable to those in non‐native species, highland native species had significantly lower elevational ranges in roadsides compared to the adjacent vegetation. We conclude that roadsides indeed change the elevational ranges of a variety of species. These changes are not limited to the expansion of non‐native species along mountain roads, but also include both upward and downward changes in ranges of native species. Roadsides may thus facilitate upward range shifts, for instance related to climate change, and they could serve as corridors to facilitate migration of alpine species between adjacent high‐elevation areas. We recommend including the effects of mountain roads in species distribution models to fine‐tune the predictions of range changes in a warming climate.     

Postglacial migration supplements climate in determining plant species ranges in Europe

The influence of dispersal limitation on species ranges remains controversial. Considering the dramatic impacts of the last glaciation in Europe, species might not have tracked climate changes through time and, as a consequence, their present-day ranges might be in disequilibrium with current climate. For 1016 European plant species, we assessed the relative importance of current climate and limited postglacial migration in determining species ranges using regression modelling and explanatory variables representing climate, and a novel species-specific hind-casting-based measure of accessibility to postglacial colonization. Climate was important for all species, while postglacial colonization also constrained the ranges of more than 50 per cent of the species. On average, climate explained five times more variation in species ranges than accessibility, but accessibility was the strongest determinant for one-sixth of the species. Accessibility was particularly important for species with limited long-distance dispersal ability, with southern glacial ranges, seed plants compared with ferns, and small-range species in southern Europe. In addition, accessibility explained one-third of the variation in species' disequilibrium with climate as measured by the realized/potential range size ratio computed with niche modelling. In conclusion, we show that although climate is the dominant broad-scale determinant of European plant species ranges, constrained dispersal plays an important supplementary role.     

Identifying the abiotic and biotic drivers behind the elevational distribution shift of a parasitic plant

• Climate change will alter the biotic and abiotic environment and dissipate ecological barriers, reorganising maps of current distribution of parasites and their hosts. In this study, we analyse the population dynamics of the parasitic plant Viscum album subsp. austriacum and explore key biotic (host availability and seed dispersal) as well as abiotic (temperature) factors influencing elevational distribution.
• The study was conducted along an elevational gradient of a Mediterranean mountain, covering the distribution belts of three potential pine hosts: Pinus halepensis (1300–1500 m), P. nigra (1300–1900 m) and P. sylvestris var. nevadensis (1600–2000 m). Along this gradient, we measured multiple variables of mistletoe population (prevalence, abundance and demographic profile) and different factors that might define the current mistletoe distribution (host suitability and availability, temperature and seed dispersal services).
• We found a decline in mistletoe prevalence and abundance with increasing elevation, detecting larger values of both variables at lower elevations of the most suitable host (Pinus nigra). Pinus sylvestris var. nevadensis was a suboptimal but suitable host for the parasite at high elevations. Mistletoe found suitable temperatures and seed dispersal services all along the gradient, being able to recruit at any site.
• With warming temperatures, the presence of suitable vectors for parasite dispersion, and the presence of a sub‐optimal host (P. sylvestris var. nevadensis) at the mountain top, mistletoe currently has a window of opportunity to expand its present geographic distribution to the summits.

     

Self-compatibility and plant invasiveness: Comparing species in native and invasive ranges

A longstanding hypothesis (“Baker's rule”) is that plant invasiveness is facilitated by floral self compatibility rather than self incompatibility. Extending this idea, invasive species whose individuals vary in degree of self compatibility within the native range might be self compatible in invading or weedy populations, due to natural selection on the mating system. We compared mating system between native and invasive ranges for two major world invasives, one annual (Echium plantagineum) and one perennial (Solanum elaeagnifolium). For an additional annual species (Centaurea solstitialis) we compared non-weedy and weedy populations in the native range. No species was strongly spontaneously self pollinating, but the degree of self compatibility after hand pollination varied dramatically. Both annuals were self incompatible in native or non-weedy populations but self compatible in invasive or weedy ones; the reverse was true for the perennial. Individuals within populations of all three species also varied in their degree of self compatibility, suggesting a basis for natural selection, and populations of the same species sharing a status (native/non-weedy, invading/weedy) varied in average self compatibility. These results support the hypothesis that differential selection of progeny during invasion can result in self-compatible populations derived from ancestrally self-incompatible ones, but that this process may be less important in perennial species, which experience multiple opportunities for sexual reproduction. Overall, however, mating system may not operate alone and its contributions to invasiveness may be conditional on other attributes of a species including physiology, morphology, and life history.     

Upward ant distribution shift corresponds with minimum,not maximum,temperature tolerance

Rapid climate change may prompt species distribution shifts upward and poleward, but species movement in itself is not sufficient to establish climate causation. Other dynamics, such as disturbance history, may prompt species distribution shifts resembling those expected from rapid climate change. Links between species distributions, regional climate trends and physiological mechanism are needed to convincingly establish climate‐induced species shifts. We examine a 38‐year shift (1974–2012) in an elevation ecotone between two closely related ant species, Aphaenogaster picea and A. rudis. Even though A. picea and A. rudis are closely related with North American distributions that sometimes overlap, they also exhibit local‐ and regional‐scale differences in temperature requirements so that A. rudis is more southerly and inhabits lower elevations whereas A. picea is more northerly and inhabits high elevations. We find considerable movement by the warm‐habitat species upward in elevation between 1974 and 2012 with A. rudis, replacing the cold‐habitat species, A. picea, along the southern edge of the Appalachian Mountain chain in north Georgia, USA. Concomitant with the distribution shifts, regional mean and maximum temperatures remain steady (1974–2012), but minimum temperatures increase. We collect individuals from the study sites and subject them to thermal tolerance testing in a controlled setting and find that maximum and minimum temperature acclimatization occurs along the elevation gradient in both species, but A. rudis consistently becomes physiologically incapacitated at minimum and maximum temperatures 2 °C higher than A. picea. These results indicate that rising minimum temperatures allow A. rudis to move upward in elevation and displace A. picea. Given that Aphaenogaster ants are the dominant woodland seed dispersers in eastern deciduous forests, and that their thermal tolerances drive distinct differences in temperature‐cued synchrony with early blooming plants, these climate responses not only impact ant‐ant interactions, but might have wide implications for ant‐plant interactions.     

Ecological ranges of plant species in the continental and ocean regions of Asian Russia

Exeplified by representatives of Poaceae of the flora of the Asian part of Russia, the ecological ranges of the plant species confined to coastal regions and of the species growing mostly within the continental regions (Inner Asia) have been compared. The comparison was made with respect to the ecological phytocenotic groups: coniferous and mixed forests, broad-leaved forests, meadows, communities of supralittoral zone, rocks, stony outcrops, and scree. Trends in the changes of the suite of indices of the ecological ranges of species that occur in the transition from continental climatic conditions of Inner Asia to monsoon climate of the Pacific coast have been revealed.     

Patterns of species ranges, speciation, and extinction

The exact nature of the relationship among species range sizes, speciation, and extinction events is not well understood. The factors that promote larger ranges, such as broad niche widths and high dispersal abilities, could increase the likelihood of encountering new habitats but also prevent local adaptation due to high gene flow. Similarly, low dispersal abilities or narrower niche widths could cause populations to be isolated, but such populations may lack advantageous mutations due to low population sizes. Here we present a large-scale, spatially explicit, individual-based model addressing the relationships between species ranges, speciation, and extinction. We followed the evolutionary dynamics of hundreds of thousands of diploid individuals for 200,000 generations. Individuals adapted to multiple resources and formed ecological species in a multidimensional trait space. These species varied in niche widths, and we observed the coexistence of generalists and specialists on a few resources. Our model shows that species ranges correlate with dispersal abilities but do not change with the strength of fitness trade-offs; however, high dispersal abilities and low resource utilization costs, which favored broad niche widths, have a strong negative effect on speciation rates. An unexpected result of our model is the strong effect of underlying resource distributions on speciation: in highly fragmented landscapes, speciation rates are reduced.     

Contrasting patterns of intraspecific trait variability in native and non-native plant species along an elevational gradient on Tenerife,Canary Islands

Background and AimsNon-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space.MethodsAlong an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species.Key ResultsWe found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species.ConclusionsWe found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.     

The push and pull of climate change causes heterogeneous shifts in avian elevational ranges

Projected effects of climate change on animal distributions primarily focus on consequences of temperature and largely ignore impacts of altered precipitation. While much evidence supports temperature‐driven range shifts, there is substantial heterogeneity in species' responses that remains poorly understood. We resampled breeding ranges of birds across three elevational transects in the Sierra Nevada Mountains, USA, that were extensively surveyed in the early 20th century. Presence–absence comparisons were made at 77 sites and occupancy models were used to separate significant range shifts from artifacts of false absences. Over the past century, rising temperature pushed species upslope while increased precipitation pulled them downslope, resulting in range shifts that were heterogeneous within species and among regions. While 84% of species shifted their elevational distribution, only 51% of upper or lower range boundary shifts were upslope. By comparison, 82% of range shifts were in a direction predicted by changes in either temperature or precipitation. Species were significantly more likely to shift elevational ranges than their ecological counterparts if they had small clutch sizes, defended all‐purpose territories, and were year‐round residents, results that were in opposition to a priori predictions from dispersal‐related hypotheses. Our results illustrate the complex interplay between species‐specific and region‐specific factors that structure patterns of breeding range change over long time periods. Future projections of increasing temperature and highly variable precipitation regimes create a strong potential for heterogeneous responses by species at range margins.     

Fifteen-year phenological plant species and meteorological trends in central Italy

The present study was carried out in a phenological garden in central Italy that contains vegetative clones of shrubs and trees common to several international phenological gardens, such as Cornus sanguinea L.; Corylus avellana L.; Ligustrum vulgare L.; Robinia pseudoacacia L.; Salix acutifolia Willd. and Sambucus nigra L. Vegetative plant growth monitoring was carried out weekly using common international keys: BBCH07, bud break and leaf unfolding; BBCH19, young unfolded leaf; BBCH91, adult leaves; BBCH93, beginning of leaf colouring. The phenological dates thus obtained provide a model of the development for these different species in relation to the 15-year period of observation (1997–2011). From a meteorological point of view, temperature and precipitation trends were studied, with the highest anomalies during the study period recorded during the first 2 months of the year (January, February). There was relative invariance in the manifestation of the open bud phase and the contemporary advance of the young open leaves phase, particularly from 2006. This was accompanied by shortening of the leaf opening period, which appeared due to more rapid spring temperature increases over the last few years. The advance tendency of the BBCH91 phase showed adult leaves from the first summer weeks with fully green foliage monitored for a long time. Generally, the autumn leaf colouring phase tended to remain constant, with the exception of Salix acutifolia and Sambucus nigra, for which, on the other hand, the first leaf development phases appeared to be most likely influenced by the photoperiod.     

Effects of altitude and climate in determining elevational plant species richness patterns: A case study from Los Tuxtlas,Mexico

Altitudinal changes of composition and richness of montane plant assemblages are complex, depending on the taxonomic group and gradient conditions, with different factors involved that are directly altitude-dependent (e.g., temperatures, air pressure) and altitude-independent (e.g., precipitation, cloud cover, area). In order to assess the relative impacts of temperature, precipitation, air humidity, and area of altitudinal belts on plant diversity, we analyzed diversity patterns of five species-rich groups, mostly herbaceous plants, in 74 forest plots along three climatically contrasting elevational transects from humid tropical lowland vegetation up to cloud forests at Los Tuxtlas, Mexico. We recorded 278 plant species, with ferns being the most species-rich group followed by orchids, bromeliads, aroids, and piperoids. The most striking results were the contrasting patterns and model results for terrestrial and epiphytic taxa. Whereas the richness of all terrestrial species taken together did not change significantly with elevation, vascular epiphytes showed increasing species numbers with altitude. However, a number of individual terrestrial taxa showed also significant elevation-related changes: aroids showed a marked decline with hight, orchids and piperoids increased, and ferns displayed a hump-shaped pattern with highest richness in mid-altitudes. Among the epiphytes, aroids declined while most other groups increased with altitude. This distinction is relevant for projections of responses of plant communities to climate change, which will lead to increased temperatures and to changing precipitation and cloud condensation regimes and thus will likely affect terrestrial and epiphytic species in different ways.